V. P. Kholodova

Selenium regulates the water status of plants exposed to drought.

The interest of researchers in selenium has sharply increased worldwide within the past two decades. Selenium is an important microelement, which is present at extremely low concentrations in bacteria, animals, and humans. Selenium is not only able to increase the general resistance of the organism to biopathogens, but also exerts a protective effect against certain oncological diseases and even immunodeficiency [1]. The protective effect of selenium in animal cells is accounted for by some proteins whose active center contains socalled selenium amino acids (selenocysteine and selenomethionine). These proteins are powerful antioxidants.

Adaptation of the Common Ice Plant to High Copper and Zinc Concentrations and Their Potential Using for Phytoremediation

A facultative halophite Mesembryanthemum crystallinum L. (the common ice plant) was shown to grow successively at the high concentrations of Cu and Zn. Although 25 µM CuSO4 or 800 µM ZnSO4 retarded markedly plant growth, they did not interfere with the completion of plant development and the formation of viable seeds. In such plants, leaves accumulated more than 200 µg of Cu and 1700 µg of Zn per 1 g of dry weight. A damaging effect of heavy metals (HMs) was manifested in a reduced content of water in leaves and proline accumulation in them. As copper is a metal with transient valence, copper salts are more toxic than zinc salts, which was manifested in a stronger inhibition of the chlorophyll synthesis. Both HMs induced oxidative stress, as evident from increased activities of guaiacol peroxidase and lipoxygenase. Moderate Cu and Zn concentrations did not damage cell membranes in leaves, as evident from the absence of their action on electrolyte leakage either under optimum conditions or after heat treatment. A capability of a substantial HM accumulation by the common ice plant and their considerable transport to shoots (up to 50 µg of Cu and 560 µg of Zn per plant) make it possible to consider the common ice plant as a promising phytoremediator.

Biological effects of high copper and zinc concentrations and their interaction in rapeseed plants

In experiments with rapeseed (Brassica napus L., cv. Westar) plants, it was confirmed that copper was considerably more toxic than zinc. The toxic effects of 50 and 150 μM CuSO4 were comparable to those of 1000 and 2500 μM ZnSO4. The analysis of the effects of these concentrations of copper and zinc on photosynthetic pigment contents and on the rate of lipid peroxidation did not reveal any reasons for different toxicities of these heavy metals (HM). Among biological effects studied, significant differences were found in the organ distribution of these metals in plants grown on both the standard medium and the medium with high concentrations of copper or zinc. Copper retained in the roots in relatively small amounts and was poorly transported over the aboveground part of the plants. It stayed mainly in the lower leaves, and its distribution changed only a little during the recovery of plants following the HM treatment. In contrast, zinc proved to be highly mobile, it was concentrated in the upper leaves and actively transported when the plants were transferred to a medium with the optimal HM concentrations. High copper concentrations slowed strongly zinc uptake by the roots but practically did not change its movement over the plant. In contrast, high zinc concentrations facilitated copper uptake by the roots but reduced its transfer to the aboveground organs. The data presented here allow us to hypothesize that biological peculiarities of organ and cellular distribution of copper and zinc in plants and interaction of these HM play an important role in the toxic effects of high concentrations of these HM and the mechanisms of adaptation to them at industrial environmental pollution used by rapeseed plants.

Current state of the problem of water relations in plants under water deficit

The review presents current literature data on the mechanisms maintaining plant water balance or those providing for tolerance to its disturbance. We consider the processes enabling the changes in the transpiration rate under water deficit due to changes in stomatal conductivity and the changes in the rate of leaf growth, as well as the role of hydraulic and hormonal (ABA, ethylene, cytokinins) signals in their regulation. Factors involved in the improvement of water use by the regulation of stomatal movements are also regarded, e.g., transcription factors, kinases, GTP-binding proteins, aquaporins participating in CO2 transfer. Negative consequences of stomata closure induced by the disturbances in gas exchange, ROS generation, and accelerated senescence and the ways of their overcoming (with the involvement of antioxidants and cytokinins as factors of senescence delay) are discussed as well. The great attention is paid to the mechanisms maintaining plant growth and transpiration under water deficit due to the optimization of water uptake (modulation of hydraulic conductivity and relative activation of root growth). It is emphasized that the role of ABA in adaptation to water deficit is not limited only to stomatal closure but also concerns the regulation of root growth and assimilate inflow to reproductive organs. Dual significance of this hormone in the growth regulation is considered: direct inhibitory and mediated stimulatory action (via normalization of water relations). The contradictory data about changes in aquaporin capacity for water transfer and their role in the changes of hydraulic conductivity under water deficit are discussed. Apparently, this contradiction may be related to specific features of water transport in various plant species (relative contribution of apoplastic and symplastic pathways) and also to the effects of such factors as an increase in the hydraulic resistance of the apoplast due to the depositions of lignin and suberin, vessel cavitation, and changes in their anatomy on hydraulic conductivity under water deficit.

Osmolyte accumulation in different rape genotypes under sodium chloride salinity

Physiological mechanisms of two rape (Brassica napus L.) genotype adaptation to chlorine salinity were investigated. The plants of two cultivars (Olga and Westar) differing in salt tolerance were grown in the pots filled with Perlite on the Hoagland and Snyder’s medium under controlled conditions. At a stage of 3–4 true leaves, the plants experienced 7-day-long salinity induced by a single addition of NaCl to the nutrient medium in order to attain desired final salt concentration (from 50 to 400 mM). The obtained results showed that a greater salt tolerance of cv. Olga plants (as compared with cv. Westar) could be accounted for by a capability of their root cells to uptake water under high salinity (300–400 mM NaCl), which is evident from a greater content of water in the tissues of cv. Olga. This was ensured by a sharp fall of the osmotic potential of the cellular contents (down to −2.3 MPa) at a low water potential of nutrient solution owing to more active uptake of Na+ (57–61 µeq/g fr wt) and K+ (210–270 µeq/g fr wt) as well as active accumulation of proline (30–50 µmol/g fr wt). The latter is caused by a reduced activity of proline dehydrogenase and retarded degradation of this osmolyte. It is important that, in contrast to less tolerant genotype, the rape plants of salt-resistant cultivar were able to maintain the K+/Na+ ratio at a rather high level at salinity of different degree, which made it possible to preserve ionic homeostasis under adverse conditions.

Effect of copper excess in environment on soybean root viability and morphology

The effects of increase copper concentrations in medium (10–150 μM CuSO4) on growth and viability of the roots of two-week-old soybean seedlings (Glycine max L., cv. Dorintsa) were studied. Copper excess suppressed biomass accumulation and linear plant growth; copper affected root growth much stronger than shoot growth. The presence of 10 μM CuSO4 in medium suppressed accumulation of plant biomass by 40% and the root length by 70%; in the presence of 25 μM CuSO4, these indices were equal to 80 and 90%, respectively. In the presence of 50 μM CuSO4, roots ceased to grow but biomass and shoot length still increased slightly. 150 μM CuSO4 was lethal for plants. The earliest sign of excessive copper toxicity was the accumulation of MDA, indicating activation of membrane lipid peroxidation. A significant increase in MDA content was observed at plant incubation in medium with 10 μM CuSO4 for 1 h; in this case, the content of copper in the roots increased from 36 ±1.8 (in control) to 48 ± 2.4 μg/g dry wt. The number of dead cells (permeable for the dye Evans Blue) was doubled in the presence of 200 μg/g dry wt within the root; this occurred in 72 h of growth in medium with 10 μM CuSO4, in 6 h at 25 μM CuSO4, in 3 h at 50 μM CuSO4, and 1 h at 150 μM CuSO4. Toxicity of copper excess was manifested stronger in dividing and elongation cells of the root apex (root meristem and the zone of elongation) than in more basal root regions. Copper excess resulted in the formation of breaks in the surface cell layers of the root tips and affect root morphology. When plant grew in medium with 10 μM CuSO4, a distance of lateral root formation zone from the root tip decreased markedly, and spherical swellings were formed on the tips of lateral roots. The higher copper concentrations (50 and 150 μM) suppressed completely the development of lateral roots.

Physiological Mechanisms of Enhancing Salt Tolerance of Oilseed Rape Plants with Brassinosteroids

The ability of brassinosteroids, such as 24-epibrassinolide (EBL) to increase the resistance of oilseed rape plants (Brassica napus L.) to salt stress (175 mM NaCl) was investigated along with the possible mechanisms of their protective action. Seedlings were grown for three weeks on the Hoagland-Snyder medium under controlled conditions. The experimental plants were treated with either (1) 175 mM NaCl, or (2) 10−10 M EBL, or (3) 175 mM NaCl plus 10−10 M EBL by adding the corresponding components to the growth medium. The exposure was 7 and 14 days. As compared to the control, salinization inhibited plant height by 33–35%, reduced leaf area by 2.0–2.5 times, reduced 2.5- and 2-fold plant fresh and dry weight, respectively, reduced water content of plant tissues by 26–31% and, twofold, the content of chlorophylls a and b. Plants responded to NaCl by developing oxidative stress conditions, lowering the osmotic potential of the cell contents down to −2 MPa, accumulating proline (by 43–52 times) and low-molecular-weight phenolics (by 1.9–2.7 times). Oilseed rape plants were shown to respond to salinization with an increase of endogenous content of steroid hormones: 24-epibrassinosteroids (24-epibrassinolide and 24-epicastasterone), 24S-methyl-brassinosteroids (brassinolide and castasterone), and 28-homobrassinosteroids (28-homobrassinolide and 28-homocastasterone); such evidence indirectly confirms the involvement of brassinosteroids in the development of salt tolerance. Adding EBL to the nutrient medium under optimal growth conditions did not significantly affect the indices under study. Under salt stress, EBL showed a pronounced protective effect: stem growth was fully restored, plant assimilation area increased by as much as 67–76% as compared to the control index, fresh and dry weight largely recovered (up to 85–92% of the control values), and the inhibitory effect of NaCl on photosynthetic pigments was diminished. Exogenous EBL impeded the development of NaCl-dependent lipid peroxidation and increased the osmotic potential of the leaf cell contents. The protective effect of EBL under salt stress was probably associated with EBL antioxidant effect, rather than the hormone-induced accumulation of proline and of low-molecula-weight phenolics, as well as with the ability to regulate water status by maintaining intracellular ion homeostasis.

Can stress-induced CAM provide for performing the developmental program in Mesembryanthemum crystallinum plants under long-term salinity?

The development of CAM-type photosynthesis is one of the adaptation mechanisms to severe water deficit. It provides plants with carbon dioxide and permits efficient water spending under extreme environments. In common ice plants, a complete switch from C3 to CAM photosynthesis was observed on the seventh day of salinity (0.5 M NaCl). The indices characterizing this switch were: (1) induction of phosphoenolpyruvate carboxylase; (2) diurnal changes in the organic acid content, which are characteristic of CAM plants, and (3) suppression of transpiration during the daytime. A decrease in the osmotic potential (ψπ) of the leaf sap, which occurred on the second day of salinity, preceded these changes. After long-term salinity stress (four–five weeks), ψπ attained extremely low values (–4.67 MPa), which made possible the water uptake by the root system. The restoration of the ψπ balance between cell compartments resulted from the accumulation of compatible solutes in the cytoplasm, proline primarily, which possesses osmoregulatory and stress-protective properties. This means that a complex of adaptive mechanisms is required for the realization of the common ice developmental program under salinity. These mechanisms maintained plant capacity to uptake water and permitted its efficient utilization. They triggered the development of stress-induced CAM-type photosynthesis, maintained the low osmotic potential in the cell sap, regulated the composition of macromolecules in the cell microenvironment, provided for water storage in tissues, and reduced the time of plant development. A comparison between the time-courses of CAM development and a decrease in the transpiration rate permitted us to suggest that a combination of low ψπ and CO2 in the leaf cells could serve as a signal for the induction of CAM-dependent gene expression in terrestrial plants.

Maize plant growth and accumulation of photosynthetic pigments at short- and long-term exposure to cadmium

A wide range of cadmium concentrations (from 4 to 200 μM for seedlings and up to 2 mM for germinating kernels) was used to assess Cd toxic effects on maize (Zea mays L.) plants at the different developmental stages: germinating kernels, seedlings (4–9 days), and juvenile plants (34 days). Cd accumulation in plant organs was followed, and its lethal concentration was elucidated. In maize, cadmium was accumulated predominantly in roots; in shoots it was mainly accumulated in the lower leaves, and the higher was leaf position the lower was Cd content in it. At high concentrations (80 and 200 μM), kernels became the substantial cadmium depot. Germinating kernels manifested the lowest sensitivity to cadmium; seedlings were more sensitive; the inhibition of juvenile plant growth attained 90% and more. In the tested range of concentrations, cadmium suppressed shoot mass accumulation harder than that of roots. In 34-day-old plants, water content in shoots was stronger reduced than in roots. Plant death was also manifested earlier in shoots. It was concluded that maize plant sensitivity to cadmium increases with plant growing and that, under conditions of normal mineral nutrition, cadmium inhibits shoot growth more severe than root growth.

Physiological mechanisms of adaptation of alloplasmic wheat hybrids to soil drought

We studied physiological mechanisms of plant adaptation to drought for two alloplasmic wheat (Triticum aestivum L.) hybrids (APHs) on the cytoplasm of rye (Secale cereale L.) or ovate goatgrass (Aegilops ovata L.) and two standard regionalized spring wheat cultivars, Kometa and Priokskaya. In response to plant tissue dehydration, APHs rapidly reduced the transpiration rate and lost much less water than standard cultivars. During drought, peroxidase activity was significantly increased only in APH on the rye cytoplasm, whereas it declined substantially in cv. Kometa. Peroxidation of lipids (POL) was activated in cv. Kometa stronger than in hybrids, which also indicates that, in this cultivar, there was no complete detoxification of hydrogen peroxide under stress conditions. After watering resumption, APHs displayed a better capacity for reparation than standard cultivars, which was manifested in peroxidase activation and POL suppression, i.e., in more complete reduction of the oxidative stress consequences. We concluded that a higher APH drought resistance, as compared with standard cultivars, was determined by their more efficient antioxidant defense and a better capacity for recovery.